Long-term changes in vegetation and soil chemistry in a calcareous and sandy semi-natural grassland

Calcicole plant species are vulnerable to acidification and fertilization, caused by deposition and changes in land use, since they are adapted to nutrient-poor calcareous conditions. In this study we used vegetation data (vascular plants, bryophytes and lichens) from 1964 and 1985 and stored soil samples from 1966 to investigate long-term soil chemistry and vegetation changes in a semi-natural, sandy calcareous grassland in southern Sweden. In the re-investigation in 2008 we found that increased decalcification due to acidification could not be verified. The plant community had changed from stress-tolerant calcareous grassland towards a community promoted by higher nutrient availability. Furthermore, the cover of species indicating calcareous conditions had decreased. A decline in the cover of species adapted to alkaline, phosphorus-poor conditions may be due to increased nutrient availability, but there were also indications that the vegetation had changed due to overgrowth by woody plants. This long-term impoverishment of the plant community highlights the need for appropriate management of calcareous grasslands, in order to limit the nutrients available in the soil and prevent overgrowth by shrubs and trees.     

Nutrient relations in calcareous grassland under elevated CO2

Plant nutrient responses to 4 years of CO₂ enrichment were investigated in situ in calcareous grassland. Beginning in year 2, plant aboveground C:N ratios were increased by 9% to 22% at elevated CO₂ (P < 0.01), depending on year. Total amounts of N removed in biomass harvests during the first 4 years were not affected by elevated CO₂ (19.9 ± 1.3 and 21.1 ± 1.3 g N m⁻² at ambient and elevated CO₂), indicating that the observed plant biomass increases were solely attained by dilution of nutrients. Total aboveground P and tissue N:P ratios also were not altered by CO₂ enrichment (12.5 ± 2 g N g⁻¹ P in both treatments). In contrast to non-legumes (>98% of community aboveground biomass), legume C/N was not reduced at elevated CO₂ and legume N:P was slightly increased. We attribute the less reduced N concentration in legumes at elevated CO₂ to the fact that virtually all legume N originated from symbiotic N₂ fixation (%N_dfa ≈ 90%), and thus legume growth was not limited by soil N. While total plant N was not affected by elevated CO₂, microbial N pools increased by +18% under CO₂ enrichment (P = 0.04) and plant available soil N decreased. Hence, there was a net increase in the overall biotic N pool, largely due increases in the microbial N pool. In order to assess the effects of legumes for ecosystem CO₂ responses and to estimate the degree to which plant growth was P-limited, two greenhouse experiments were conducted, using firstly undisturbed grassland monoliths from the field site, and secondly designed `microcosm' communities on natural soil. Half the microcosms were planted with legumes and half were planted without. Both monoliths and microcosms were exposed to elevated CO₂ and P fertilization in a factored design. After two seasons, plant N pools in both unfertilized monoliths and microcosm communities were unaffected by CO₂ enrichment, similar to what was found in the field. However, when P was added total plant N pools increased at elevated CO₂. This community-level effect originated almost solely from legume stimulation. The results suggest a complex interaction between atmospheric CO₂ concentrations, N and P supply. Overall ecosystem productivity is N-limited, whereas CO₂ effects on legume growth and their N₂ fixation are limited by P. Received: 12 July 1997 / Accepted: 15 April 1998     

放牧对草原植物功能性状影响研究进展

植物功能性状的表达和植被环境适应性相关,植物功能性状之间的权衡变化体现了植物在放牧胁迫下资源的重新整合和获取.本文总结了放牧干扰下植物功能性状表达的差异性,着重将放牧干扰与植物功能性状相结合,介绍了植物功能性状的变异来源是植物遗传特征与环境过滤相互协调的结果,归纳了放牧对植物营养性状、繁殖性状的影响,以及植物可以通过调...     

养分与水分添加对荒漠草地植物钠猪毛菜功能性状的影响

分析水分与养分添加对植物功能性状的影响, 对揭示植物对环境变化的响应和适应规律至关重要。该文通过施氮与增水(包括增雨、增雪)共6种处理(对照(N₀W₀)、雨添加(N₀W₁)、雪添加(N₀W₂)、氮添加(N₁W₀)、氮雨添加(N₁W₁)、氮雪添加(N₁W₂)), 对荒漠草地进行氮肥添加、水分添加的实验。实验持续了3年, 第4年以荒漠草地建群种植物钠猪毛菜(Salsola nitraria)植物功能性状为研究对象, 量化分析了钠猪毛菜10种植物功能性状对氮素和水分添加的不同响应。得出以下结论: (1)双因素方差分析结果表明, 施氮、增水的交互作用对钠猪毛菜的茎鲜质量、叶鲜质量、叶面积、比叶面积、叶饱和含水量、叶干物质含量存在显著影响(p < 0.05), 而施氮主效应对钠猪毛菜各植物功能性状指标不存在显著影响、增水主效应对钠猪毛菜各植物功能性状指标不存在显著影响。(2)在钠猪毛菜10种植物功能性状指标中, 株高、茎鲜质量、茎干质量、叶饱和鲜质量、叶干质量、叶面积、比叶面积的最大值均出现于N₁W₂处理中(p < 0.05), 其次为N₀W₀处理。叶饱和含水量的最大值出现于N₀W₀中, 其次为N₁W₂处理中。N₁W₀处理显著降低了叶饱和含水量(p < 0.05); 叶干物质含量值在各处理下均高于N₀W₀, 其中N₀W₁处理的叶干物质含量显著高于N₀W₀ (p < 0.05)。(3)各处理下, 叶干物质含量与比叶面积为负相关关系。在准噶尔荒漠草地, 钠猪毛菜植物功能性状在不同处理下的不同表现是适应环境变化的结果。     

养分添加对退化草地豆科植物草木犀功能性状的影响

养分亏缺是造成草地生态系统退化的重要因素, 养分添加被认为是促进退化草地恢复的重要手段。豆科植物作为草地生态系统的重要功能种群, 其对养分添加的响应尚不清楚。该研究选取典型豆科植物草木犀(Melilotus officinalis)为研究对象, 在浑善达克沙地开展了鸡粪添加(L0-L6分别为0、500、1 000、1 500、2 000、2 500和3 000 kg·hm_-2·a^-1)控制实验, 探究了养分添加对豆科植物功能性状的影响。鸡粪添加对草木犀的叶片氮(N)含量、磷(P)含量、钾(K)含量、最大净光合速率(P_nmax)、水分利用效率(WUE)、株高(H)和比叶面积(SLA)均有显著影响。随鸡粪添加量的升高, 草木犀叶片N、P、K含量均线性升高, 当添加量达到3 000 kg·hm_-2·a^-1 (L6)时, N、P、K含量升高幅度分别达到15%、67%和25%; 草木犀的H、SLA、P_nmax和WUE先上升后下降, 转折点出现在鸡粪添加量为1 500 kg·hm^-2·a^-1 (L3)时, 以8月份为例, L3较L0的升高幅度分别达到51%、18%、12%和38%, 而L6与L0则无显著差异。因此, 适量的鸡粪添加能显著提高豆科植物的光合能力和水分利用能力, 改善其生长状况, 有利于豆科植物种群的维持。     

Large-scale manipulation of plant litter and fertilizer in a managed successional temperate grassland

Plant litter may play an important role in herbaceous plant communities by limiting primary production and influencing plant species richness. However, it is not known how the effect of litter interacts with fertilization. We tested for the role of litter and fertilization in a large-scale experiment to investigate effects on diversity and biomass of plant species, growth forms, native vs. non-native groups, and abiotic ecosystem components (e.g., soil moisture, PAR). We manipulated plant litter (removed vs. left in situ) and nutrient availability (NPK-fertilized vs. unfertilized) for 4 years in 314-m² plots, replicated six times, in an old-field grassland. While many of our species-level results supported previously published studies and theory, our plant group results generally did not. Specifically, grass species richness and forb biomass was not affected by either fertilization or plant litter. Moreover, plant litter removal significantly increased non-native plant species richness. Relative to native plant species, all of our experimental manipulations significantly increased both the biomass and the species richness of non-native plant species. Thus, this grassland system was sensitive to management treatments through the facilitation of non-native plant species. We coupled biotic and abiotic components within a nonmetric multidimensional scaling (NMS) analysis to investigate treatment effects, which revealed that specific treatments altered ecosystem development. These results suggest that fertilization and plant litter may have larger impacts on plant communities and on ecosystem properties than previously understood, underscoring the need for larger-scale and longer-term experiments.     

Vegetation and seed bank in a calcareous grassland restored from a Pinus forest

Question: What is the importance of the seed bank in the maintenance of the restoration potential of a 60‐year‐old abandoned calcareous grassland overgrown by Pinus trees? Location: ‘Les Pairées’, province of Luxembourg, Belgium. Methods: The seed bank and the above‐ground vegetation were surveyed in three adjacent stands, previously forming a unique calcareous grassland: a 60‐year‐old Pinus forest, a four‐year‐old clear‐cutting and a typical calcareous grassland. Floristic diversity was compared among stands and between vegetation and seed bank. Results: The species richness of the vegetation and the seed bank was significantly lower in the Pinus forest. More floristic similarities were found between the clear‐cutting and the calcareous grassland. Seed bank was essentially transient, dominated by annual species. Its correspondence with the above‐ground vegetation was weak. Conclusion: Very few calcareous grassland species have persisted in the Pinus stand. Four years after clear‐cutting, the stand was nearing restoration towards a calcareous grassland. Seed longevity in the soil was not the most explicative factor. Dispersal of propagules from adjacent sources was also important.     

Vegetation dynamics in a coastal grassland of Hawaii

Vegetation development following goat removal in a tropical grassland with a Mediterranean rainfall seasonality on leeward Hawaii is reported from two permanent monitoring sites. The two sites are equipped with experimental exclosures against feral goats. The sites were monitored annually for a decade (1971–80) for changes in cover by species. The vegetation changes reported relate to both inside- and outside-exclosure developments.Four dynamic categories were recognized among the grassland species following goat removal. These are the decreasers, the increasers and the persistent species. The latter were divided into stable and oscillating persisters. Attention was drawn to the oscillating persisters, as exemplified by the endemic vine Canavalia kauensis, and to the absence of any correlation of its oscillation pattern to the variation in year-to-year rainfall. It was concluded that the oscillation pattern resides in the population it self and can be attributed to its life-cycle phases. It is suggested that synchronized life-cycle dependent death in local populations is an important dynamic phenomenon in certain plant communities. This phenomenon may be considered as separate from phenology and succession, but of similar significance in understanding the processes of vegetation dynamics.Nomenclature of plant names follows St. John (1973).     

短期增温和降水减少对半干旱沙质草地群落特征的影响

【目的】为探究短期增温和降水减少对沙质草地植物群落物种多样性、功能性状及生产力的影响,进一步揭示沙质草地植物群落特征对气候变化的响应。【方法】以科尔沁沙质草地植物群落为研究对象,利用开顶式生长室(OTC)模拟增温,研究降水减少0%、20%、40%和60%与增温的交互作用对沙质草地群落组成、物种多样性和功能性状的影响及群落特征与环境因子关系。【结果】结果显示:（1）短期增温和降水减少使沙质草地植物群落组成和物种优势度发生变化,其中优势物种猪毛蒿和达乌里胡枝子在短期增温和降水减少处理下均明显降低,短期增温和降水减少均显著降低了植物群落盖度和地上生物量。（2）短期增温显著降低了Margalef丰富度指数、Simpson指数、Shannon多样性指数和Pielou均匀度指数,短期降水减少对沙质草地供试4种物种多样性指标均无显著影响。（3）短期增温显著提高了植物群落高度和叶干物质含量,显著降低叶厚度,短期降水减少显著提高了叶干物质含量和叶厚度,显著降低了植株高度和比叶面积。（4）短期增温是影响沙质草地物种多样性和功能性状的主要环境因素。【结论】综上可知,短期增温和降水减少改变了沙质草地群落特征,温度是影响沙质草地群落特征的主要环境因子。     

Plant community dynamics in a calcareous grassland under climate change manipulations

This study investigates the effects of field manipulations of local climate to determine the potential impact of climate change on plant community dynamics in a calcareous grassland. The experimental site is located in a grassland at the Wytham estate, Oxfordshire, UK. The one hectare study area is within a 10 ha abandoned arable field on Jurassic corallian limestone. Two climate change scenarios were used: warmer winters with increased summer rainfall and warmer winters with summer drought. Plant cover and species richness were significantly increased in plots receiving supplemented summer rainfall, while the amount of litter was significantly reduced. Litter formation was significantly increased by winter warming and drought. The responses of the plant community to the climate manipulations were related to the life-history attributes of the dominant species. Seedling recruitment was limited by microsite availability, which also varied in the different climate manipulations. The results are discussed in terms of successional dynamics. They suggest that warmer winters may delay succession, as gap formation in the sward will provide sites for colonisation of annuals, thereby enabling their persistence in the sward. Under wetter conditions during summer, perennial grasses tend to close the sward, thereby inhibiting the establishment of later successional species.     

退化草地物种种内和种间功能性状变异对放牧的响应

研究草地植物功能性状变异对放牧干扰的响应机制,有助于解析草地生态系统群落构建和功能多样性维持机制及植物对环境的适应及权衡策略。以科尔沁沙地退化草地为对象,研究围封和放牧草地物种多种功能性状(植株高度、根系长度、叶面积、根面积、叶片干物质含量、地上干物质含量、根系干物质含量、比叶面积、比根长和根冠比)变异特征及其对放牧干扰的响应机制。结果表明: 退化草地物种种间性状变异明显高于种内变异,种间性状变异对总体变异的贡献率占比高达70.2%～95.1%,而种内变异仅为4.9%～29.8%,但群落构建中物种的种内变异仍不可忽视。放牧草地物种种间性状变异低于围封草地,且放牧草地种内变异增加而种间变异减小。放牧导致不耐牧的优良禾本科牧草叶面积和叶片干物质含量下降而比根长增加,但耐牧的杂类草则通过增加叶面积和叶片干物质含量并降低比根长来提高在群落中的优势地位。退化草地对放牧响应较为敏感的功能性状有叶面积、叶片干物质含量、比根长和比叶面积。植物叶片性状和根系性状自身及彼此间均呈显著正相关,但放牧会增强根系性状的协同效应而减弱叶片性状的协同效应。说明放牧会驱动植物个体和种群功能性状权衡策略发生改变,进而起到调控植物群落结构和功能的作用。     

Nitrogen mineralisation and plant nitrogen acquisition in a nitrogen-limited calcareous grassland

A field study measured the rate of soil mineral N supply and its effects on plant biomass and N accumulation in a 13-year-old, naturally regenerating, calcareous grassland. Gross rates of N mineralisation (2 μg g⁻¹ day⁻¹, i.e. 0.69 kg ha⁻¹ day⁻¹), assessed using ¹⁵N pool dilution, were at the lower end of the range previously reported for grasslands. Weekly additions of liquid N fertiliser ([NH₄]₂SO₄, NH₄NO₃ or KNO₃) and, to a lesser extent the addition of water, increased plant growth substantially, demonstrating that the primary constraint to plant growth was low N availability. In plants that had received NO₃⁻, the activity of the inducible enzyme nitrate reductase in shoots initially increased in proportion to the amount of NO₃⁻ supplied. However, as above-ground herbage accumulated, nitrate reductase activity declined to similar low levels in all treatments, despite the continuance of the constant NO₃⁻ additions. The decline in NR specific activity reflected declining tissue NO₃⁻ concentrations, although total plant NRA may have remained constant during the period of study. The study has shown that plant growth is limited by low N mineralisation rates and indeed the soil is a sink for much added N. Low water availability provides an additional constraint on N mineralisation in this calcareous grassland soil. Any disturbances in the N cycle which increase the availability of mineral N will result in a substantial increase in plant growth within this ecosystem.     

Plant traits and functional types in response to reduced disturbance in a semi‐natural grassland

Abstract. Question: How do functional types respond to contrasting levels of herbage use in temperate and fertile grasslands? Location: Central France (3°1’E, 45°43’N), 870 m a.s.l. Methods: Community structure and the traits of dominant plant species were evaluated after 12 years of contrasted grazing and mowing regimes in a grazing trial, comparing three levels of herbage use (high, medium and low). Results and Conclusions: Of 22 measured traits (including leaf traits, shoot morphology and composition, phenology), seven were significantly affected by the herbage use treatment. A decline in herbage use reduced individual leaf mass, specific leaf area and shoot digestibility, but increased leaf C and dry matter contents. Plants were taller, produced larger seeds and flowered later under low than high herbage use. Nine plant functional response types were identified by multivariate optimization analysis; they were based on four optimal traits: leaf dry matter content, individual leaf area, mature plant height and time of flowering. In the high‐use plots, two short and early flowering types were co‐dominant, one competitive, grazing‐tolerant and moderately grazing‐avoiding, and one grazing‐avoiding but not ‐tolerant. Low‐use plots were dominated by one type, neither hardly grazing‐avoiding nor grazing‐tolerant, but strongly competitive for light.     

Fifty years vegetation development of a xerothermic calcareous grassland in Central Europe after heavy disturbance

Fifty years ago on the south-facing slope of a limestone hill 4 km NE of Göttingen (Germany) a small plot of 4 m² size was chosen in a xerothermic calcareous grassland (Mesobromion alliance) for investigation. The formerly grazed slope was abandoned a few years prior to the study, but in 1987 intermittent grazing started again. The mean annual temperature in Göttingen (1953–2003) was 8.8 °C on the average; the annual mean increased by 1.1 °C during the study period. Mean annual precipitation amounted to 642 mm (393–1093 mm). The mean monthly precipitation during summer (April–August) varied in the range between 29 and 101 mm. In the years 1953–2003 the presence of species was recorded annually in 400 subplots (100 cm² each). In part of this time also the amount of bare soil within the plot was recorded, and standing crop and the nitrogen contents in leaves of Bromus erectus were measured in the immediate surrounding of the plot.In fall 1953 the vegetation was removed completely in half of the plot, in the other half the two dominant grasses Bromus erectus and Brachypodium pinnatum were left. The differences between the two halves vanished already after 2 years (starting phase 1953–1955). Thus, only results concerning the plot as a whole are reported here. During the investigation three periods could be distinguished. After the starting phase in period I (1955–1986) total presence, presence of the most important species, species richness, number of endangered species, diversity and evenness increased. In this way the grassland regenerated after the heavy disturbance, and a dense, species-rich vegetation formed. Only a few species showed maximum values already within the first 10 years and faded out later.Period II (1987–1997) spanned 10 years of grazing. During this time total presence, presence of many important species, species richness, number of endangered species, diversity and evenness decreased, whereas areas with bare soil increased. Only two species showed maximum values during this phase, and just one new species invaded (Lolium perenne). During this period a more open type of grassland was formed. Period III (1998–2003) spanned 5 years without grazing and only one grazing year (2002). Here, a partial regeneration of a more closed grassland took place with traits similar to period I.Several phenomena occurred continuously through all three periods. Bromus erectus always was the dominating species. Nevertheless, proportions of graminoids (in % total presence) were lowered, and the proportion of herbs increased. The ratio Bromus erectus/Brachypodium pinnatum was determined mainly by Brachypodium. Seedlings of woody plants were always present, but in most cases survived only one summer. Shrubs did not settle inside the plot, although small bushes developed in the immediate surrounding. In summary the 50-year process can be described as a slow recovery of a calcareous grassland after very heavy disturbance, modified by intermittent grazing, without formation of shrub vegetation.     

The competition–dispersal trade‐off exists in forbs but not in graminoids: A case study from multispecies alpine grassland communities

Much theoretical evidence has demonstrated that a trade‐off between competitive and dispersal ability plays an important role in facilitating species coexistence. However, experimental evidence from natural communities is still rare. Here, we tested the competition–dispersal trade‐off hypothesis in an alpine grassland in the Tianshan Mountains, Xinjiang, China, by quantifying competitive and dispersal ability using a combination of 4 plant traits (seed mass, ramet mass, height, and dispersal mode). Our results show that the competition–dispersal trade‐off exists in the alpine grassland community and that this pattern was primarily demonstrated by forbs. The results suggest that most forb species are constrained to be either good competitors or good dispersers but not both, while there was no significant trade‐off between competitive and dispersal ability for most graminoids. This might occur because graminoids undergo clonal reproduction, which allows them to find more benign microenvironments, forage for nutrients across a large area and store resources in clonal structures, and they are thus not strictly limited by the particular resources at our study site. To the best of our knowledge, this is the first time the CD trade‐off has been tested for plants across the whole life cycle in a natural multispecies plant community, and more comprehensive studies are still needed to explore the underlying mechanisms and the linkage between the CD trade‐off and community composition.     

Species-area relations in a New Zealand tussock grassland,with implications for nature reserve design and for community structure

Abstract. A tussock grassland, in Blackrock Reserve, New Zealand, was sampled thoroughly at scales ranging from 0.01 m x 0.01 m to 20 m x 20 m, to investigate species-area relations of relevance to plant community structure, and to offer a pointer to reserve design. In total, 96 species were found. Of the native vascular species among these, 20% were new records for the reserve. For the total and the native vascular flora the observed points fell midway between the Arrhenius and Gleason fitted curves. Cryptogams fitted the Arrhenius model well, with z close to the 0.26 value expected for isolates under Preston's Canonical hypothesis. Extrapolation of the Arrhenius curve to the whole of the New Zealand uplands gave values far too large; the Gleason curve gave values much too low. A General Root model is introduced. It fits the observed species richnesses at various quadrat sizes considerably better than previous models. When biogeographic limitations on species richness are included, extrapolation of the General Root curve accurately predicts the size of the upland flora. This fit, the lack of an asymptote, and the rarity structure, are compatible with a random and individualistic model of community structure. However, there are differences in species-area relation between vascular and cryptogamic plants, which cautions against expecting any universal type of community structure. Extrapolations using the General Root model suggest that if the ideal is a 10 km x 10 km reserve, a reserve one tenth that size would contain 81% of the native species in that ideal, and the present Blackrock Reserve contains 67%.